Botanica Marina Vol. 45, 2002, pp.


© 2002 by Walter de Gruyter · Berlin · New York

A Revision of Amansia glomerata C. Agardh, Amansia rhodantha (Harvey) J. Agardh and Melanamansia glomerata (C. Agardh) R. E. Norris (Rhodophyta: Rhodomelaceae)
A. D. R. N’Yeurt
Jeune Equipe Terre Océan, Université de Polynésie Française, BP 6570, Faa’a Aéroport, Tahiti 98702, French Polynesia,

Species of Amansia and Melanamansia from the tropical Pacific region have been re-examined, and compared with material from the Indian Ocean, Japan, Taiwan and the Philippines. All specimens examined from the Pacific region (except Hawaii and some from New Caledonia and Fiji) lack pseudo-pericentral cells, thus placing them in Amansia. Melanamansia glomerata is only reported from Hawaii, New Caledonia, Fiji, the Philippines,Vietnam, Malaysia and for the first time in the Indian Ocean from Kenya.Type material of Amansia rhodantha (Harvey) J. Agardh from Mauritius was compared and found not significantly different from the Pacific Amansia material (including type and isotype material of Amansia paloloensis South et Skelton from Samoa, which is reduced to synonymy), thus extending the distribution range of A. rhodantha as a pantropical entity.

The genus Amansia was erected by Lamouroux (1809: 322) and is placed in the tribe Amansieae of the family Rhodomelaceae (Hommersand 1963: 335). The tribe is characterised by dorsi-ventral, erect thalli arising in tufts from a common basal disc. Most genera have five pericentral cells around the axial cell (Figs 1, 2) except Kuetzingia Sonder and Protokuetzingia Falkenberg, which have six pericentral cells (Kylin 1956). The two first-formed pericentral cells lie on the dorsal side of the characteristically inrolled apex of the mature lanceolate blades. In Amansia the dorsal and dorsi-lateral pericentral cells further divide to form immediately behind the apex a distromatic wing or ala, whose cells are arranged in alternate layers, with secondary lateral pit-connections (Fritsch 1945: 570). Recently, Norris (1988b, 1995) re-examined species of Amansia in the Indian Ocean and Hawaii, and discovered that some members of this genus differed consistently by having the first and second pericentral cells each producing a lateral pseudo-pericentral cell lying adjacent, but not connected to, the axial cell (Norris 1988b: 214; Fig. 3). Species previously ascribed to Amansia that had these pseudo-pericentral cells were transferred to a new genus (Melanamansia R. E. Norris 1988b: 217). Notably, Hawaiian plants previously ascribed to the common tropical species Amansia glomerata C. Agardh (whose type locality is Hawaii) were transferred to Melanamansia glomerata (C. Agardh) R. E. Norris, while Indian Ocean ‘Amansia glomerata’ were put under the earlier name Amansia rhodantha (Harvey) J. Agardh based

on their lack of pseudo-pericentral cells (Norris 1995). The genus Amansia currently consists of four species, the Caribbean A. multifida Lamouroux (the type of the generic name), A. loriformis R. E. Norris and A. rhodantha (Harvey) J. Agardh from the Indian Ocean and Malaysia, and A. paloloensis South et Skelton from Samoa. Melanamansia currently consists of twelve species, M. daemelii (Sonder) R. E. Norris, M. glomerata (C. Agardh) R. E. Norris, M. japonica (Holmes) R. E. Norris, M. mamillaris (Lamouroux ex C. Agardh) R. E. Norris, M. pinnatifida (Harvey) R. E. Norris, M. pumila (Sonder) R. E. Norris, M. scalpellata (Tanaka) R. E. Norris, M. seagriefii R. E. Norris, M. serrata (Harvey) R. E. Norris and M. mitsuii (Tanaka) Yoshida. Following these revisions by R.E.Norris,several authors (Yoshida et al. 1995,N’Yeurt et al. 1996,Payri and N’Yeurt 1997, Millar 1999, Millar et al. 1999) made the nomenclatural change from Amansia glomerata to Melanamansia glomerata in their checklists and herbarium collections from the tropical Pacific, but until recently no detailed studies verifying these changes as correct have been done. In 1995, samples of ‘Amansia glomerata’ from Rotuma Island (north of Fiji) were sent to Dr R.E.Norris,who confirmed (pers. com.) that pseudo-pericentral cells were present, and on the basis of this assumed finding the name Melanamansia glomerata was used in later publications (N’Yeurt 1996,N’Yeurt et al. 1996).Later,in the course of on-going investigations on the Samoan algal flora, South and Skelton (1999) found a species of Amansia lacking pseudo-pericentral cells, which they described as new (A. paloloensis South et Skelton), while in Malaysia and Vietnam (Masuda et al. 2000, Masuda

Tahiti. leg. 1997. in particular paying attention to important anatomical features such as the presence or absence of pseudo-pericentral cells and the number of segments between lateral veins of the blade (Norris 1995). SUVA-A 2402 (Lectotype) and liquid-preserved isotypes 5479L (tetrasporic). leg. – French Polynesia: Moorea. 11 Jul. ps = pseudo-pericentral cells). Photomicrographs were obtained using an Olympus BX-50 compound microscope. Y. Enomoto (SAP). 3897). 1998. Punaauia. 5480L (cystocarpic). 9 Feb. 2 Jul. R. Results Representative material examined Amansia rhodantha – Japan: Hanasado Beach. M.Agardh to co-exist in the flora. Fig. Sections were made using a freezing microtome. a = axial cell. leg. Fig. 1998. 18 Sep. leg. A. 2. 6 Jun. Okinoerabujima Island (60 km north of Okinawa) 27 Sept. Okinoerabujima Island (60 km north of Okinawa). SUVA-A 5436L). Payri (8/84. Arrangement of pericentral cells in species of Amansia and Melanamansia (modified from Norris 1988b. leg. 3899). leg. showing intrusion of the first derivative of the first-formed pericentral cell into the axial ring (Suva. leg. N’Yeurt (UPF 347). 3896. and stained with cotton blue in lactophenol or 1% acidified aniline blue in 60% clear corn syrup. – Samoa: Palolo Deep Reserve. 1998. 3892. Rarotonga. Skelton. 13 May 1982. Furusato. Okidomari. SUVA-A 5438L). A. 3893. 5481L (spermatangial)].. Tanegashima Island. Taharaa. Kagoshima: 9 August 1959. New Caledonia. P. leg. 1999. Baba (SAP 056163). N’Yeurt (UPF 584). 1984. in EPHE). Skelton. 1998. 3895. D. E. A. Material and Methods Specimens were collected by reef-walking. M. deep algal rim. Skelton and Dave [SUVA-A 1565 (holotype). The results of these investigations are described below. 19 Mar. Tahiti. Melanamansia glomerata. leg. A. developed and printed in the laboratory. N’Yeurt In order to clarify the taxonomy of species hitherto ascribed to Melanamansia glomerata. Tombleson and T. it was decided to revise records in the light of recent findings. snorkelling or SCUBA diving. Amansia rhodantha. 1. leg. Afiti and D. Amansia rhodantha and Amansia glomerata. com.‘Al’ and ‘CG’ refer to specimens housed at IRD. C. cross section of the lectotype from the Hawaiian Islands (LD 42600) showing the first two dorsal pericentral cells each developing a pseudo-pericentral cell (ps) that lies adjacent to the axial cell. otherwise as in Holmgren et al. M. and a camera-lucida attachment was used to make anatomical drawings. a detailed morphological and taxonomic study on some selected members of the Amansieae (Wilson and Kraft 2000) echoed the opinion that the genus Amansia in Australia was in need of critical revision. U. Macrophotography was done with a Nikon F2A in conjunction with Kodak Plus-X pan 125 film. Prepared slides were examined using a Zeiss compound microscope. Fiji. Amansia rhodantha (Beqa.) recent studies found both Melanamansia glomerata and Amansia rhodantha (Harvey) J. Figs 1–3. Faletose (SUVA-A 3894. Fig. – Taiwan: Anon. 3898. SUVA-A 1566 (isotype). P. Fiji. (1981). 1998. A. A. P. 3. pers.232 A. N’Yeurt (SUVA-A 5128). A. Sakai and S. Rintels (SUVA-A 3891. Palolo Deep. 2 Jul. . Baba (SAP 056160). Nouméa. – Cook Islands: Ngatangiia. and stored in 5% buffered Formalin in seawater prior to shipment to the laboratory. P. 25 October 1984 (SAP 046978). leg. Skelton and Dave (SUVA-A 1567). Most recently. 1989. 1–5 = order of formation of pericentral cells.

. leg. B. Vatualailai. leg. 1985. 1876.Dumaguete City.d. Cribb (SAP 051714).). Daida (US 086755). Garrigue (SUVAA 5436L). Namada. anon.Solong-On.. Negros Oriental. Nanuyalevu. Table I presents the average results of morphological measurements carried out on Amansia rhodantha. Taxonomic results Amansia rhodantha (Harvey) J. 17 Jul. A. – Lord Howe Island: 18 Aug. (MURU. 21/22-Sept. leg. SIPHILEXP-78 (US 088534). Snellius-II (L 0108601). n. 14 Feb. 16 Nov. J. J. A. A. C. M de Robillard (L 0108599). South (SUVA-A 5431). Farlow (L 0108597). P. (US 02890). Bay of Sanggar. Snellius-II (L 0108602). Magnesia Virae. leg. J. Montebello Islands.. South (SUVA-A 2460). Hapmak. SIPHILEXP-78 (US 013760). 25 Feb. 06 Sept. leg. Eagles Nest. P. Oahu. Garrigue (CG 77c. Suva Barrier Reef. Skelton and G. 1988.3 2. South (SUVA-A 5544. 1819. leg. Naukathura Island.7 54. leg. Carlson (SUVA-A 281). leg.Negros Oriental. 18 March 1992. leg. Silva et al.1 22. 17 Mar. Villeneuve (SUVA-A 1737). Viti Levu.d. Hildebrandt (L 0108600). Wilkes Exploring Expedition (US 04057a). – Fiji: ‘Feejee’. 84 (L 0108603). Species Thallus height (cm) Blade width (mm) Blade length (mm) Serrations (1 –3)* Cortication (1 –3)* Rosettes (1 –3)* Amansia rhodantha Amansia paloloensis Melanamansia glomerata 39. 1069). Figs 4–10 . LD 42600 / 0772). Potipot Is.11 Feb. Jul. – Australia: Heron Island QLD.leg. 3 and 15 Oct.P. 27 Dec. R. leg. Banilad. 1982. Kasahara (SUVA-A 1013. Bahati. Dumaguete. – Mauritius: Delesseria rhodantha Harvey (holotype) (= Amansia rhodantha): Cap Malheureux. J. R. JH240). leg. Huisman (MURU. leg. paloloensis and Melanamansia glomerata specimens examined in this study.. Ovalau. H. 3 and 5 March 1992. n. 1965. Yasawa Group. 1980. 10 July 1992. SIPHILEXP-78 (US 088531). Hanauma Bay Beach Park. – Kenya: Mombassa. Yasawa Islands. leg. R127). Salayer. Inhaca Peninsula. N’Yeurt (SUVA-A 5437L). Viti Levu.Siaton. SIPHILEXP-78 (US 088530).2 53. J.Palawan. 1838–42. leg. 9 Sept. W. SIPHILEXP-79 (US 088533). 1996: 472. leg. Huisman (MURU. Average values of selected characters for Amansia paloloensis. R. 1985. leg. Amansia rhodantha and Melanamansia glomerata surveyed in this study. Hakula. leg.W. E. H.9 11. Richer de Forges (AL 288d-CG 77a). G.d. A. W.23 May 1978. 1995. C. Lectotype.Cuyo. Melanamansia glomerata – Hawaiian Islands: Hawaiian Islands. leg. B. 1984. Keats. R. Agardh). 1997.7 2 2 1 2 2 2 2 2 2 * As measured on an index of (1 = little) to (3 = pronounced) – Rotuma: Hoféa. 30 Jun.. 84.Malo. 3 Feb. W. 1979. – Mozambique: Ponta Abril.Siquijor Is. B262). leg. Candelaria Zambales. J. leg. NR 248).6 3. Oahu. Oahu.14 May 1979. 16 Oct. 26 Nov. leg. C. leg. 1954. M. Isaac (L 0108594). N’Yeurt (SUVA-A 346. 24 April 1996. leg. Jurien Bay WA. P. leg. S.. A. 1876. 3 Jun 1978. – Fiji: ‘Feejee’. leg. H. leg. Ballou (SUVA-A 5438L). E. S. R. South in lit. 1067). 1995. leg. fide G. 6 Jun 1978. Tantabiddi. leg. Chesterfield. Cordero and Tuangpalan (US 091395). Littler (US 163490). 1982. CG 77b). 1981. 28 Nov. Kasahara (SUVA-A 1012. n. 10 Aug.Canipo Is. Wilkes Exploring Expedition (US 04057b) (doubtful record. leg. – Philippines:Gao-oa. leg. leg. Barrow Island. 07 Oct. 1834). Beqa Lagoon. L 0108598). 29 May 1978. M. 14 March 1998. 2000: 188. Agardh 1841: 26. 5 June 1967.Revision of some Amansia and Melanamansia species 233 Table I. Lopta. C. c. Cordero and Tuangpalan (US 091394). 13 July 1972. Viti Levu. 1973.. 1980. SIPHILEXP-78 (US 088532). leg. Tangangge. Ningaloo. D. Cannon (SAP 052062). G. leg. figs 1–11 (as Amansia glomerata C. 1994 (SUVA-A 797). 10 May 1978. – New Caledonia: Lajeu Sudouest. Villeneuve (SUVA-A 1774). – Kenya: Mombassa. leg. G..d. Kipukai. Ti Tree. Unia. Amansia rhodantha: n.Moreland (US 50851). in TCD (Telfair. – New Caledonia: Lajeu Sudouest. 1984. leg. Rukuruku Village. G. leg. Plongeurs IRD (AL 288a. Catanduanes. A. figs 37–42. 15 Oct. E. n. leg.Negros Oriental. W. Huisman (CWC 1997. Negros Oriental. 1995. 24 Nov. Kauai. 84. 16 March 1991. AL 288b). Masuda et al. Basay. leg. Barrow Island. Hildebrandt (L 0108596. Garrigue (CH1). M. J.d.5 3. Suva Reef. Gaudichaud (‘Amansia glomerata’. B. leg. Huisman. Isaac (L 0108595). 25 September 1999. Norris 1988b: 211. 1979.9 18. 1838–42. leg.Ilocos Norte. Lawai Kai. Diani Beach. see text). B79). Huisman (MURU. leg. leg. leg. Mauritius.leg. A.Calindagan Reef. leg. M. Magruder (US 086459). A.Pagodpud. 21 Sep. 347). leg. Sumbawa. leg. – Philippines: Actin. 1983. Kauai. – Indonesia: Cape Batu Kerapo. Jul.

showing both lax and dense growth forms. Mauritius. Fig.234 A. Fig. D. South Pacific (scale bar = 10 µm). Figs 4. Fig. 9. in TCD (scale bar = 25 µm). in TCD (scale bar = 200 µm). in TCD (scale bar = 2. showing axial cell (a) surrounded by five pericentral cells. Axial structure of blade. . 8. and a pair of budding branchlets from the midrib in the lower half. Type material of Delesseria rhodantha Harvey. Surface view of segment cells of blade alae. R. 5. Amansia rhodantha. 10.5 mm). Secondary branchlet. showing marginal serrations. Rehydrated type material of Delesseria rhodantha Harvey. N’Yeurt Figs 4–10. 6. Fig. Detail of tetrasporangial stichidia on lateral endogenous branch. 7. Type material of Delesseria rhodantha Harvey. SUVA-A 5437L. US 02890 (scale bars = 3 mm). from Rotuma Island. Indian Ocean. Habit of mature plant from the type locality.

showing both lax branching and the presence of rosettes (arrowheads) (scale bar = 4 mm). Fig. Fig. Fig. 12. 11. 4. Tetrasporangial stichidia of Amansia rhodantha from Beqa. showing lateral endogenous branches and midrib.5 mm). . Amansia paloloensis from Samoa.Revision of some Amansia and Melanamansia species 235 Figs 11–16. Fig. SUVA-A 5436L (scale bar = 50 µm). 15. Compare with Fig. Detail of blades. Detail of morphologically distinct mature rosette and corticated axis. SUVA-A 5479L (scale bar = 1 mm). Habit of mature plant. SUVA-A 3892 (scale bar = 12 mm). Fiji. Segment cells of blade alae in surface view. Fig. 13. Holotype of Amansia paloloensis. 16. 14. Fig. SUVA-A 3892 (scale bar = 2. Detail of the holotype (SUVA-A 1565). Note marginal cystocarps (arrowheads). SUVA-A 1565 (scale bar = 25 µm). with numerous rosette axes.

becoming narrower and disappearing towards the apex. 24. Fig. on average 53 mm high. and two displaced rows of regularly arranged hexagonal alar cells 20–29 × 92–129 µm in surface view. 6. with in-rolled leaf apex. showing pseudo-pericentral cells (arrowheads) surrounding (but not directly connected to) axial cell (a). pl. N’Yeurt et al. Norris 1995: 66) is on average 4. with more or less pronounced marginal serrations 1–4 mm long (which represent endogenous branches) always present. AL288b (scale bar = 5 mm). 1995: 150. Philippines. they are up to 4 cells long and non-vesiculate. New Caledonia. The number of segments between alternating veins (determinate branches covered by the corticating pericentral cell derivatives. 436B. Fritsch 1945: 570. 19. 18. figs 127. Isaac and Isaac 1968: 24. up to 100 (on average 40) mm high. figs 2–14 (type locality: Palolo Deep Marine Reserve. pl. fig. up to 750 µm long and 250 µm broad. and are crisp and lanceolate. Blade margins are smooth. 128. Apia. Agardh 1824: 194. Fig. CG77a (scale bar = 1 mm). Dublin). Distribution: Tropical Indian Ocean. – Melanamansia glomerata (C. fig. Tetrasporangial stichidia of plant from Oahu. 197. fig. Philippines. 1999: 573. Synonymy: Rytiphlaea rhodantha (Harvey) Decaisne 1842: 358. colourless and uniseriate dorsal trichoblasts up to 8 cells long develop on every second or third axial cell (or on every segment in determinate marginal branchlets). 9). Secondary blades develop as successive pairs on the dorsal side of the midrib. 185. figs 19–21. SUVA-A 5431 (scale bar = 50 µm). Detail of blade of plant from Mombassa. Yamada and Tanaka 1938: 86. Branching is lax to profuse and usually forms deep-red leafy rosettes of secondary branchlets when fully mature. The stem of plants is thick and cartilaginous. with a central axial cell 28–30 µm in diameter surrounded by five pericentral cells 36–40 µm in diameter (Figs 1. Kenya. Payri and Meinesz 1985: 511. Blades of plant assumed to be from the Fiji Islands (see text). New Caledonia. Fig. Kenya. 212a.8–14. and are initially vesiculate. A distinct midrib up to 1 mm wide is present in the lower half of ultimate branchlets. Agardh) R. in up to 10 regularly disposed pairs. and are composed of lanceolate to ovate blades to 100 mm long and 3 mm wide. Known distribution: Hawaiian Islands. with a range of 1–5. fig. up to 35 mm long and 6 mm broad. Tetrasporangia occur in flattened. E. East Africa. Millar et al. Falkenberg 1901: 416. Plants are brownish-red. Axial structure of plant from Lajeu. with a range of 3–6. Weber-van Bosse 1923: 369. Deciduous. 1. fig. Fig. 267. showing smooth margins and elongate shape. Fig.4. Agardh 1824: 247. Samoa). L 0108598 (scale bar = 5 mm). holotype in TCD. composed of two overlapping layers of elongate hexagonal cells in transverse rows. surrounded by five pericentral cells. South-East Asia.8 mm in diameter. Abbott 1999: 404. 19). with the first two dorsal pericentral cells each developing a smaller pseudo-pericentral cell 13. Plant from Mombassa. Norris 1995: 66) is usually 4. Carposporangia are lanceolate. Garrigue and Tsuda 1988: 63. Mauritius. . 0. 6E (unverified record). E. Tsuda and Wray 1977: 103.5 µm in diameter that lies adjacent to the axial cell (Figs 3. Lewis and Norris 1987: 22. Kenya. Marginal teeth are up to 1 mm long and 445 mm broad. L 0108598 (scale bar = 700 µm). pl. 22. Fig. US 4057b (scale bar = 1 mm).236 A. Internal structure is cellular. to 0. 4B. Kylin 1956: 544. 1987: 61. pl. Isaac 1956: 188. Individual tetrasporangia are spherical and tetrahedrally divided. 20. 21. XXV. 100–112 µm in diameter. N’Yeurt Basionym: Delesseria rhodantha Harvey 1834: 151–152. The number of segments between alternating veins (determinate branches covered by the corticating pericentral cell derivatives. Norris 1988b: 211. Ultimate branchlets are ecorticate. Millar 1999: 523 (unverified record). Cribb 1983: 106. US 091395 (scale bar = 5 mm). 77–110 µm thick. Plant from Unia. A central midrib is present. figs 1–3 (type locality: Cap Malheureux. figs 1–11. Amansia paloloensis South et Skelton 1999: 247. Jaasund 1976: 131. Note that the stichidia is twice laterally branched. containing up to 10 Figs 17–24. figs 14–29. The thallus is rose-red. 1996: 87. Procarps and curved spermatangial heads are 200–450 µm in diameter and are dorsal on the involucral tips of endogenous lateral branches. denuded below. 118C–J. Payri and N’Yeurt 1997: 897. Japan. N’Yeurt 1996: 428. Plant from Potipot Island. or with regular serrations (endogenous branches). R. Fiji. stem-like and rigid in basal portions. New Caledonia. 17. and the apex is characteristically in-rolled. New Caledonia. Fig. Silva et al. 20. pinnately arranged marginal stichidia up to 340 µm long and 290 µm in diameter. 2. Figs 11–16 Misapplied names: Amansia glomerata C.5–0. while cystocarps 700–1000 µm in diameter occur usually singly and terminally on endogenous branches. Norris 1995: 67. fig. plate CXXVI. Coppejans and Millar 2000: 333 (unverified record). figs 1–13. Norris 1995: 67. 1. Womersley and Bailey 1970: 336. each cell is about 20–20 × 87–100 µm in surface view. Yoshida et al. D. e–i. 31 fig. 7 mm broad. Agardh) R. Blade of plant from Lajeu. Tetrasporangial stichidia are pinnately arranged on either side of blade margins. CG77a (scale bar = 10 µm). Melanamansia glomerata (C. 198. 85–110 µm thick. pl. Trichoblasts are sparse. The main axis is irregularly branched. Fig. Figs 17–24 Basionym: Amansia glomerata C. Okamura 1900: 71. Tropical Pacific Ocean. The central axial cell is 28–30 µm in diameter. Melanamansia glomerata. 23. 25–42 × 83–125 µm. Vietnam.

procarps and cystocarps occur on endogenous marginal branches. the colour of the Melanamansia plants was noticeably brownish-red. . Spermatangial heads are curved and arranged in marginal clusters. and stained the herbarium mounting paper. In all cases.Revision of some Amansia and Melanamansia species 237 pairs of tetrahedrally-divided tetrasporangia which are 50–100 µm in diameter.

New Caledonia. Exploring Expedition. From this study. rhodantha from the Indian Ocean and other localities. more coriaceous branchlets with less apparent midribs (this study). generally smaller. Fig. com. Characters such as the degree of rosette formation. and Ceramium. one of which (US 04057a) is Amansia rhodantha. 3) which distinguishes that plant from A. rhodantha from the type collection and localities in Mauritius. when pseudo-pericentral cells were found to be absent) are considered conspecific with Amansia rhodantha. fig. isotype and other collections of A. However. rhodantha. R. On the other hand. 7) shows features similar to those seen in blades of South Pacific material previously ascribed to Amansia glomerata. while Melanamansia becomes dark brown. Melanamansia glomerata in the tropical Pacific Wilson and Kraft (2000: 366) commented that the generic segregation of Melanamansia from Amansia is probably not warranted. it appears to be conspecific with the latter species and thus extends the status of A. plate CXXVI. the Philippines. the absence of blades organized in morphologically distinct tight axes around the parent axis. Amansia tends to bleach to translucent white. thicker. since the presence of pseudo-pericentral cells is an inconsistent attribute in species of Neurymenia. the Cook Islands and French Polynesia (Table I) with A. with for instance A. paloloensis from the type locality. Japan. paloloensis from A. Fiji. The habit and description of the holotype by Harvey (1834: 151.13). 12). serrate branches and lax ‘rosettes’ of secondary blades issued from the weakly developed midrib. 2) indeed shows a plant with loriform. a comparison of material of A. 23) but the plant has not been collected in Fiji waters since. rhodantha as described and illustrated by Norris (1988b: 211). Agardh from Hawaii (LD 42600 / 0772) clearly shows the presence of paired dorsal pseudo-pericentral cells in a blade cross section (Fig. degree of cortication of the midrib and position of the reproductive stichidia are equally quite variable in plants from the same and similar populations. paloloensis. Rotuma. length of the blades. additional distinguishing characteristics of Melanamansia are a darkly coloured brown thallus which stains the mounting paper when dried (possibly linked to differences in chemical composition. and a weakly developed midrib as characters distinguishing A. and these authors considered the lack of highly corticated axes. having tight rosettes as well as very lax. The status of Amansia paloloensis South et Skelton South and Skelton (1999) described a new species of Amansia from Samoa. D. The present author has re-examined the holotype. an additional field test to distinguish Amansia spp. and found a significant number of thalli with morphologically distinct primary corticated axes and relatively tight and lax rosettes of secondary blades on rosette axes (Figs 11. Kuetzingia.). Eastern Australia. These plants also had blades with well-developed midribs in the lower half (Fig. The Wilkes collection consists of two plants. N’Yeurt Discussion The holotype of Delesseria rhodantha Harvey An examination by the author of a rehydrated blade fragment of the holotype of Delesseria rhodantha Harvey in TCD (Fig. Smith (1979: 38–40) points out that the accounting and . and trichoblasts lacking protective vesicles when young (Norris 1988b). apparently because the brown pigments (presumably polyphenols) are less susceptible to the bleaching action of the preservative (as is the case in species of brown algae such as Sargassum and Turbinaria). Vidalia. paloloensis showed no major difference in the range of characteristics among them. On the basis of these observations. from Melanamansia glomerata was found: when preserved in 4% Formalin-seawater and exposed to ambient light for several days. rhodantha (South and Skelton 1999: 249). Mozambique and Western Australia. loriform blades reminiscent of some specimens of A. In the Fiji Islands. a re-examination of the lectotype of Amansia glomerata C. material from the Tropical Pacific and Indian Oceans previously known as Amansia glomerata (or misidentified as Melanamansia glomerata. is comparable to A. Kenya. a single record for this genus exists (collected from ‘Feejee’ by the Charles Wilkes U. Masuda pers.238 A. Now that the Samoan species has been compared with genuine herbarium material of A.S. More importantly. rhodantha as a pan-tropical entity. figs 1–4) indicate features quite similar to those found in the Amansia species considered in this study from localities in both the Indian Ocean and the tropical Pacific Ocean. The reproductive and trichoblast anatomy of A. While there is no doubt as to the generic identity of the latter specimen. indicating that the morphology of the species is influenced perhaps by environmental factors. M. between 1838 and 1842 (US 04057b. and absence of pseudopericentral cells. rhodantha from Western Australia (MURU B262. which is welldocumented and illustrated in the protologue of the species. The drawing of the holotype (South and Skelton 1999: 246. and a cross section of the blade clearly shows the five pericentral cells around the axial ring. while the other (US 04057b) is Melanamansia glomerata. paloloensis. MURU 248). 14) and corticated rosette axes (Fig. as well as Amansia species from Taiwan. including the presence of both tight and lax rosettes of secondary branches and alar cells on the primary axes. regularly alternate marginal serrations which are not as pronounced as in Amansia rhodantha and which are often absent.

In most instances. except the Hawaiian samples. Proliferations from the ventral midrib (aside from secondary branchlets) were only found to be abundant in some of the Indonesian Amansia rhodantha specimens from Salayer (e. fig. type of endogenous branching. Masuda. pers. and would seem to function as attachment haptera. position of the reproductive axes and the distinctness of the axial ring in rosette axes (Norris 1988a. the Philippines. however it is clear that the degree of rosette formation and cortication of the axes is not a reliable taxonomic character at the species level in Amansia. 19). Generally.e. There is a possibility it might have been mixed with collections from neighbouring localities such as the Solomon Islands. when present). Herbarium and liquid-preserved samples of Amansia and Melanamansia species from the Pacific. Ecological studies would be needed to verify these hypotheses. in some cases. with the blades smooth and superficially reminiscent of those of some Sargassum species (Figs 22–24). When pseudo-pericentral cells were present. which was the main reason for separating the species from A. while younger plants have less cortication and more lax or absent rosettes of blades. Also. and the presence of tight and lax rosettes of secondary branches on the same plant (Figs 11–13). Indonesian and Indian Ocean region were sectioned and examined for pseudo-pericentral cells. The illustration by Harvey of the holotype of Delesseria rhodantha (1834. the vesiculate (in Amansia) or non-vesiculate (in Melanamansia) nature of trichoblasts. These ventral. an examination of the holotype and liquid-preserved and pressed isotypes of A. Wilson and Kraft 2000). which is a characteristic feature of both Amansia and Melanamansia (Norris 1988b) can vary from lax to tight. both on the same plant and within individuals in similar populations. both have an ala) and are arranged in ‘lax’ rosettes (South and Skelton 1999: 247. calm waters (such as blue holes. 1) shows both the presence of tight and lax rosettes of secondary branches. 2). 5.). Melanamansia glomerata is unlikely to be confused with other Melanamansia species because of its unique combination of characteristics (Norris 1988b. it was observed that rosettes of blades along the secondary axes. 41) similarly report the absence of pseudo-pericentral cells in A. irregularly dichotomous proliferations lack alar cells.g. perhaps an adaptation to the exposed habitats where the plants came from. the description of the species states that branches have the same shape as the main axis (i. However. and a main axis with alar cells not very distinct from secondary branches. rhodantha. Validity of current criteria to distinguish the Amansieae Characters that have proved useful in distinguishing species in the Amansieae include the presence or absence of pseudo-pericentral cells. while in Malaysia Masuda et al. Fiji and New Caledonia (Table I). in the case of Amansia paloloensis). thus placing them in Amansia. so it cannot be confirmed at this stage if M. From an examination of the specimens at hand. The presence or absence of these pseudo-pericentral cells hence appears to be a good generic criterion. pl. and some samples from Kenya. Fijian and French Polynesian specimens previously ascribed to Melanamansia (N’Yeurt et al. M. they were clearly apparent and about half the size of pericentral cells (Fig. and in some instances lacking altogether. com. Herbarium specimens of ‘Amansia glomerata’ from southern Japan housed in SAP (Hokkaido University) were examined. some specimens were virtually indistinguishable from specimens of A. older plants tend to have thicker. fig. in the case of A. CXXVI. pseudo-pericentral cells were universally absent. alternate marginal proliferations (which represent non-determinate or endogenous secondary branchlets) 2–5 mm long (Fig. The author has unsuccessfully tried to obtain material from the latter locality. more corticated secondary and primary axes with tight rosettes. paloloensis revealed a wide range in the degree of rosette formations and cortication of the secondary axes and presence of alae on primary axes. more exposed conditions (such as on fringing reefs) favour tight rosette formation (which is more hydrodynamically resistant) while more lax arrangements are possible in deeper. the presence or absence of bladelets in rosettes. 1995). Norris (1988b: 211) reported that rosette axes in .Revision of some Amansia and Melanamansia species 239 recording of the Wilkes Expedition collections was not very satisfactory. and hence the precise locality and date of this Melanamansia collection from Fiji is unknown. In Melanamansia glomerata these marginal serrations were much less pronounced (but more regular than in Amansia rhodantha. (2000: 188. in New Caledonia and Kenya). 13). Payri and N’Yeurt 1997) were reexamined. which also had a consistently thicker thallus. 7). It would seem that perhaps shallow. rhodantha.g. fig. rhodantha from Mauritius and Western Australia (compare Figs 4. and is backed by other morphological and biochemical differences (Norris 1988b. For instance. 1996. Indian Ocean Amansia rhodantha also lacked such cells. However. M. degree of marginal serrations and thallus flattening. adding to the confusion between the two genera. and found to lack pseudo-pericentral cells. glomerata occurs in the Solomon Islands. environmental factors seem to play a role in determining the degree of rosette formation. All Amansia rhodantha specimens examined had fairly prominent. L 0108603). Indeed. paloloensis. and found to consistently lack pseudo-pericentral cells and a similar observation was made on samples from Taiwan. glomerata occurs in the same habitat as Amansia rhodantha (e.

15). as the axial ring was still discernible in all sections made. but the range is reportedly less than in Amansia rhodantha. in that they are typically rhodomelaceous with pairs of tetrasporangia in up to ten rows. An examination of these elegant cells in species of Amansia (Figs 10. paloloensis specimens studied. 16) and Melanamansia show no particular difference. smaller and narrower blades. showing the currently known distribution. 20). Melanamansia glomerata from all localities studied have consistently smoother. with most specimens having segment lengths of about 100 µm. rhodantha and A. An additional potentially useful character is the number of segments between lateral ‘veins’ on the blades.). but can also occur apically. Biogeographical map for Amansia rhodantha (circle) and Melanamansia glomerata (triangle). 25. At this stage there seems to be no largely significant difference in the number of segments between alternating veins in the specimens of A. rhodantha specimens observed in this study. only being more or less surrounded by cortication. usually 3 or 4 with an average of 3. N’Yeurt Amansia from Natal have an ill-defined axial row of cells. paloloensis. Cells of the blade alae in both Amansia and Melanamansia are regularly elongate-hexagonal. These differences reinforce the conspecificity of the Amansia rhodantha and A.3. Hence the distinctness of the axial ring was not found a reliable taxonomic character among the species examined. Cover cells usually totally cover the developing sporangia. One feature which was noted in Melanamansia glomerata is that individual tetrasporangial stichidia are often laterally branched once or twice (Fig. blade dimensions. cortication of axes and rosette formation between specimens of Amansia rhodantha studied. B. as well as between A. the number ranged from 3 to 7. which was also recorded by Norris (1988b) for Hawaiian material. In A. Norris (1995: 66) counted the number of segment cells between these alternating veins for several species of Melanamansia and Amansia. which was not seen in Amansia rhodantha (Figs 6.4 (Norris 1995).240 A. D. and occur in two overlapping and slightly offset layers radiating on either side of the midrib. with the length of segments ranging from 80 to 120 µm. and found them relatively constant. as well as the generic distinctness of Melanamansia glomerata. In Melanamansia it averages 4. From Table I. as well as more corticated axes. R. com. rhodantha studied. There appears to be no notable difference in the range of thallus size. which are in fact determinate branches covered by the pericentral cell derivatives. in elongate flattened and pinnate stichidia which are usually on lateral marginal branchlets. . This also varied according to the degree of rosette formation and age of the plant. However. Tetrasporangial stichidia are basically similar in both Amansia rhodantha and Melanamansia glomerata. Fig. a number of preliminary conclusions can be made: A. This feature was examined on a range of specimens of Amansia from the South Pacific and the Indian Ocean but no significant difference was seen in these specimens. These can be easily detected in the unrolled apical portions of the blades by the developing trichoblast terminating each such alternating ‘vein’ (Norris 1995 and pers. it was noted that in Melanamansia the range was more restricted.

Marine red algae from the North coast of Papua New Guinea. W. 1976. 10 figs. T. Marine botany of the Kenya coast. East Africa Nat. M. I. Bot. 3. K. Lund. M. First edition. Nat. Bot. Utrecht. M. . Cal. Accepted 17 January 2002. Index Herbariorum. Part 1. Rhodophyceae. Acknowledgements The author sincerely thanks the following persons and institutions for the generous loan of type material and specimens used in this study: Dr John Huisman (MURU – Murdoch University). E. Part I. iii + 159 pp. (2000) confirm that the genus Amansia does exist in the Pacific Ocean. Decaisne. Notice of a collection of algae. and F. and A. Mar. pls v–vii. Skelton (SUVA – The University of the South Pacific). F. V. Essais sur une classification des algues et des polypiers calcifères de Lamouroux. II. Lord Howe Island (G. W. Norris. Tsuda. Sci. 2000. 1956. 1824. Handbook No 2. et description de cinq nouveaux genres de cette famille. ments rose-red. Mr Barrett Brooks and Dr John Sims (US – Smithsonian Institution. C. Pub. Isaac. Robin South is also thanked for his helpful comments on an earlier draft of this manuscript. Paris 1: 330–333. com. 71 pls. N. Coppejans. W. in the Mauritius. and it appears that Melanamansia glomerata is so far restricted to Hawaii (Norris 1995). Phang. Mar. and possibly other localities which are not yet adequately investigated. 2. R. 1988. K. Catalogue of marine benthic algae from New Caledonia. Taxonomic notes on marine algae from Malaysia. H. Observations sur la physiologie des algues marines. and each genus is consistently differentiated from the other by a number of anatomical and biochemical characters. trichoblasts on every second or third axial cell. South pers. initially vesiculate and composed of up to 8 cells: Amansia rhodantha. Professor Willem Prud’homme van Reine (L – Rijksherbarium. General account of the environment. Marine Red Algae of the Hawaiian Islands. Berlin. J. Bohn. 27: 7–28. Professor G. Shimada. Sapporo). Die Rhodomelaceen des Golfes von Neapel und der angrenzenden Meeres-Abschnitte. Agardh. trichoblasts non-vesiculate. communicated to Dr. Systema algarum.). Hist. Masuda. Gleerups Förlag. Marine algae of Inhaca Island and of the Inhaca Peninsula.. C. P. 1963. Norris is thanked for his helpful insights and suggestions for this research. Kawaguchi and S. and J. com. Die Gattungen der Rhodophyceen. I. Phaeophyceae. 1983.. 1956. Lewis. pigments reddish-brown. Cribb. – Absence of paired dorsal pericentral cells. Honolulu. S. xvi + 477 + [1] pp. Seven species of Rhodophyceae. E. P. Foreword. University of Tromsø. It is clear that both A. Sci. Washington). 29: 1–38. 2000. rhodantha and Melanamansia glomerata are more widely distributed than previously thought. 24 pls. and R. M. Ms Susanna Riebe (LD – Botanical Museum. with descriptions of some new and little known species. Sci. Monographie 26.A. Soc. Schofield. sparse. 35: 165–366. Falkenberg. Bot. flora and vegetation. Dr Claude Payri (UPF – Tahiti). ser. Hommersand. J. 43: 315–346. Nouv. E. Myxophyceae. J. A. Masuda pers. 1809. from “Cap Malheureux”. Mar. Kylin. xiv + 939 pp. Kenya (this study). F. Jaasund. 458 figs. 43: 181–190. E. 1945. Intertidal Seaweeds in Tanzania. J. Garrigue.A. xxxviii + 312 pp. Dr John Parnell (TCD – Trinity College. Lund). Marine Algae of the Southern Great Barrier Reef. composed of up to 4 cells: Melanamansia glomerata. J. Australian Coral Reef Society: Brisbane. cxxvi. Fritsch. 2 folded maps. Mus. Robin South and Mr Posa A. Revised Key to the Species of Amansia and Melanamansia in the South Pacific – Presence of paired dorsal pericentral cells.Revision of some Amansia and Melanamansia species 241 Conclusions The results of this study and those of South and Skelton (1999) and of Masuda et al. 17: 297–380. 1968. In historiam algarum symbolae. Smiths. Millar. 1999. pig- References Abbott. Linnaea 15: 1–50. 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